The First Described Nests of Black-Bellied Cuckoo Piaya

Johan Ingels & Mathias Fernandez 280 Bull. B.O.C. 2014 134(4)

The frst described nests of Black-bellied Cuckoo Piaya melanogaster, from French Guiana

by Johan Ingels & Mathias Fernandez

Received 9 July 2014

Summary.—Although widespread throughout Amazonia, litle is known of the breeding biology of Black-bellied Cuckoo Piaya melanogaster and its nest has not been described. We found two nests in French Guiana, both with two nestlings. The open, cup-shaped nests were constructed with twigs, pieces of vine and dead leaves within dense tangles of vines. The nestlings were mainly fed hairy caterpillars and, to a lesser extent, other arthropods. Our observations suggest that the species’ breeding biology is similar to that of the closely related and beter- known Squirrel Cuckoo P. cayana.

The non-parasitic cuckoos of the genus Piaya, Squirrel Cuckoo P. cayana and Black- bellied Cuckoo P. melanogaster, are widespread over most of the New World tropics. They possess discreet behaviour, moving furtively through the forest canopy or other tall vegetation, and are easily overlooked. They feed on a variety of arthropods, most frequently on hairy, noxious caterpillars. Of the two Piaya, only the breeding biology of the Squirrel Cuckoo is more or less known. Its open cup-shaped nest of twigs and sticks is constructed in a tangle of lianas or vines, in dense shrubbery or in overgrown trees. Clutch size is two, rarely three, white eggs (Payne 1997, 2005, Erritøe et al. 2012). Black-bellied Cuckoo occurs in eastern Venezuela, south-eastern Colombia, the Guianas, eastern Ecuador, eastern Peru, northern Bolivia, and from Pará, Amapá and across Brazilian Amazonia to northern Mato Grosso. It mainly occupies the canopy and subcanopy of terra frme forest and low forest on sandy soil, although it is occasionally observed at forest borders and in shrubby vegetation away from forest, or in savanna woodland. It is generally thought to be uncommon, but its secretive behaviour makes it difcult to observe, and the species is the least known of the Piaya cuckoos, while its nest remains undescribed (Payne 1997, 2005, Erritøe et al. 2012). In French Guiana, the species is confned to the country’s interior forests, where it is rather common, and it is absent from the litoral (Tostain et al. 1992). Only one breeding record for Black-bellied Cuckoo is mentioned in the literature. In late July 1989, adults atending a nest with nestlings were observed along the piste de Saint-Elie (c.05°17’N, 53°03’W) in French Guiana, but no details are known (Tostain et al. 1992). We report here on two nests of Black-bellied Cuckoo also found in French Guiana.

Methods The site of the two nests was an islet forming part of the Saut Mapaou, a rapid on the Approuague River (04°12’N, 52°18’W). The distance between the island and the riverbank on one side was c.10 m. On this bank, MF owns a clearing of c.0.5 ha with a shelter, which he visits at irregular intervals. The islet has a surface area of approximately 10 × 5 m and is completely covered by vegetation, with a tall tree, c.12 m high. The frst nest was constructed in the tree and was observed for a total of approximately 12 hours from a distance of c.3 m by MF between c.07.00 h and 12.00 h during the mornings of 16 and 17 August, and between c.14.00 h and 16.00 h during the afternoon of 16 August 2013. The

© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club Johan Ingels & Mathias Fernandez 281 Bull. B.O.C. 2014 134(4) second nest was constructed in the same tree, but was only visited briefy on 11 November 2013 to check the contents.

Results On 15 August 2013, the atention of MF was drawn by Black-bellied Cuckoos fying to and fro between the riverbank and the islet in the rapid. On searching the islet, MF discovered the birds’ open, cup-shaped nest c.8 m above ground in a dense tangle of vines around a fork at the tip of a horizontal branch (Fig. 1). It was constructed of twigs, pieces of dry vine and dead leaves, and resembled suspended liter, which made it difcult to locate the nest from below. The external diameter and the total height of the nest were c.25 cm and 10 cm, respectively. It contained two nestlings, approximately 8 ± 2 days old (Fig. 2). The adult cuckoos were not shy. They continued their provisioning activities in a normal way when MF was siting in the tree. By day, they brought food to the nest approximately every 45 minutes. However, from sunrise (at c.06.20 h) to c.09.00 h and from c.17.00 h to sunset around 18.30 h, provisioning was much more frequent, with intervals between feedings of just 10–15 minutes. All foraging occurred in the canopy of trees on the mainland. An adult with prey always arrived on the islet and then at the nest in the same way. On arriving at the riverbank, it frst perched for several minutes in the vegetation, observing the surroundings before fying to the islet (Fernandez & Ingels 2014). There, the adult hopped up through the vines to the nest. It always arrived from below, never from above, and always via the same route. On departing the nest, the adult few of directly or hopped to a nearby vine and then few to the riverbank. Just prior to arriving at the nest, an adult with prey would uter a few soft calls to which the nestlings would respond with begging calls. These loud begging calls could be heard on the riverbank i.e. from a distance of c.12 m. Transfer of prey and occasional removal of a faecal sac occupied <1 minute, following which the adult immediately departed. In the frst hour or so after dawn, when feeding rates were most intense, faecal sacs were removed during each provisioning, three during the frst morning and four the second morning. During the day when provisioning was infrequent, faecal sacs were removed less often. The inside of the gape of the nestlings was bright red and the palate showed small white knobs (Fig. 3). The nestlings were mainly fed hairy caterpillars and, to a lesser extent, with grasshoppers, crickets and other arthropods (Fernandez & Ingels 2014). Caterpillars brought to the nest included Automeris illustris (Fig. 4), A. hamata and A. liberia, also a Dirphia sp. probably tarquinia (Hemileucinae, Saturniidae) and Amphonyx duponchel (Sphinginae, Sphingidae). Other prey included grasshoppers, crickets (Gryllidae), bush crickets (katydids, probably Pseudophyllinae, Tetigoniidae) and cicadas (Cicadidae), as well as wandering spiders, probably a Phoneutria sp. (Ctenidae). All caterpillars of the family Saturniidae have stinging hairs, and spiders of the family Ctenidae are poisonous (F. Bénélux pers. comm.). However, we assume that the adults squeezed the possibly toxic leaf remains from the guts of caterpillars before they were fed to the nestlings (Fig. 5). On 18 August, in the late afternoon, the nest was found to be have been destroyed and the nestlings had disappeared. It was most probably predated. The loud begging calls of the nestlings and / or MF’s presence on the islet could have atracted the atention of predators. Possible predators seen in the clearing and on the islet included Micrastur forest falcons and Tayras Eira barbara. On 11 November 2013, MF observed the cuckoos again fying to and fro between the riverbank and the islet. On visiting the islet, MF discovered that they had a new nest with nestlings. It was similar to the frst and constructed in a very dense tangle of vines in the

Figure 1. Black-bellied Cuckoo Piaya melanogaster at nest within dense tangle of vines, French Guiana, August 2013 (Mathias Fernandez) Figure 2. Nestlings of Black-bellied Cuckoo Piaya melanogaster, approximately 8 ± 2 days old, French Guiana, August 2013 (Mathias Fernandez)

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Figure 3. Nestlings of Black-bellied Cuckoo Piaya melanogaster utering begging calls as an adult arrives with prey, the white knobs on reddish palate clearly visible; French Guiana, August 2013 (Mathias Fernandez) Figure 4. Adult Black-bellied Cuckoo Piaya melanogaster with noxious Automeris illustris caterpillar, French Guiana, August 2013 (Mathias Fernandez) Figure 5. Adult Black-bellied Cuckoo Piaya melanogaster with an Automeris caterpillar that has a drop of gut contents hanging from its rear, French Guiana, August 2013 (Mathias Fernandez) same tree, but 4 m lower, at a height of c.4 m. The tangle of vines was so dense that it was impossible to see or photograph the young in the nest. However, when the adults arrived with food, MF heard begging calls of two diferent young and, from the intensity and a

© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club Johan Ingels & Mathias Fernandez 284 Bull. B.O.C. 2014 134(4) comparison of these calls with the begging calls of the young at the frst nest, he estimated that they were c.1 week old. Unfortunately, the survival of this nest could not be followed.

Discussion The nest sites of Black-bellied Cuckoos described here are similar to those chosen by Squirrel Cuckoos, i.e. dense shrubbery or a tangle of lianas or vines in trees (Payne 2005, Erritøe et al. 2012). Both nests of Black-bellied Cuckoo were open cups, similar to the shallow platform or open-cup nests of Squirrel Cuckoo (Haverschmidt & Mees 1994, Payne 2005). The frst Black-bellied Cuckoo nest held two nestlings, which corresponds to the usual clutch size of Squirrel Cuckoos (Penard & Penard 1910, Payne 2005, Erritøe et al. 2012). When found on 15 August, the nestlings were estimated to be 8 ± 2 days old. With an estimated incubation period of 18–19 days as for the similar-sized Squirrel Cuckoo (Erritøe et al. 2012), eggs would have been laid around 20 July. The nestlings in the second nest were probably one week old on 11 November, which suggests that eggs were laid on c.15 October, 8.5 weeks after the frst nest disappeared and 12.5 weeks after the frst clutch was laid. Black-bellied Cuckoos probably nest several times in a season, like Squirrel Cuckoos (Payne 1997), meaning that the second clutch should not necessarily be viewed as a replacement. Our observation of approximately one feeding of the nestlings per 45 minutes during the day corresponds well with the single feeding per hour given for Squirrel Cuckoos (Skutch 1966). Many of the caterpillars and arthropods fed to the nestlings have stinging hairs and/or are noxious. The adults probably squeeze the possibly toxic leaf remains from the guts of most caterpillars, but it did not appear that stinging hairs are removed. We did not observe if the nestlings regurgitated these hairs in pellets, as adults do (Payne 1997). The white knobs on the reddish palate of the nestlings are remarkable. Skutch (1966) stated that the gape of Squirrel Cuckoo nestlings is bright red, although he did not mention the presence of white knobs or markings. The gapes of nestlings of Guira Cuckoo Guira guira, Crotophaga species and some Coccyzus species also possess bright, whitish markings that contrast strongly with the red palate (Sick 1993, Payne 1997). Such markings may assist the adults to place food in the chick’s open bill, especially in a dark nest within dense tangles of vegetation, as is often the case for Piaya cuckoos. However, the question then remains why these markings should only occur in nest-building cuckoos and not in those that parasitize host species with a domed or closed nest, e.g. Striped Cuckoo Tapera naevia and both Dromococcyx (Payne 1997). In Guyana, Squirrel Cuckoos have been recorded breeding in April and July– September (Erritøe et al. 2012), and in Surinam in July (Ribot 2013), while eggs in the Penard collection were collected in March–July and September (Haverschmidt & Mees 1994). In French Guiana, nests under construction were found in March and August, and a nest with young in July (Tostain et al. 1992). In Amapá and Pará (Brazil), nesting has been observed in January, March, May, July and August (Payne 2005, Erritøe et al. 2012). In French Guiana, Black-bellied Cuckoo nests with nestlings have been found in July (Tostain et al. 1992), and August and November (this paper). Thus, over the Guiana Shield (French Guiana, Surinam, Guyana and adjacent regions of Brazil), Piaya cuckoos appear to breed from January to November, i.e. from the onset of the short dry season (February–April), through the long rainy season (May–mid August) until the end of the long dry season (mid August–November) (Penard & Penard 1910, Payne 2005, Erritøe et al. 2012). As Squirrel Cuckoos nest several times over the course of a single season (Payne 1997) and there does

© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club Johan Ingels & Mathias Fernandez 285 Bull. B.O.C. 2014 134(4) not appear to be any seasonality in breeding by either Piaya, they probably breed year- round in this region.

Acknowledgements We thank Frederik P. Brammer, Guy Kirwan, Jan Hein Ribot, Tom Schulenberg, Arie Spaans and Paul Van Gasse for help with literature, and Marco Gaiani and José Clavijo A. of the Museo del Instituto de Zoología (Maracay, Venezuela), and Frédéric Bénélux, for identifying food items brought to the nestlings. We are grateful to Frederik P. Brammer, Olivier Claessens, Des Jackson, Guy Kirwan and Chris Sharpe for their comments and for improving drafts of this paper.

References: Erritøe, J., Mann, C. F., Brammer, F. P. & Fuller, R. A. 2012. Cuckoos of the world. Christopher Helm, London. Fernandez, M. & Ingels, J. 2014. Photospot: A nest of Black-bellied Cuckoo Piaya melanogaster in French Guiana. Neotrop. Birding 15: 63–65. Haverschmidt, F. & Mees, G. F. 1994. Birds of Suriname. Vaco Uitgeversmaatschappij, Paramaribo. Payne, R. B. 1997. Family Cuculidae (cuckoos). Pp. 508–607 in del Hoyo, J., Elliot, A. & Sargatal, J. (eds.) Handbook of the birds of the world, vol. 4. Lynx Edicions, Barcelona. Payne, R. B. 2005. The cuckoos. Oxford Univ. Press. Penard, F. P. & Penard, A. P. 1910. De Vogels van Guyana (Suriname, Cayenne en Demerara), vol. 2. F. P. Penard, Paramaribo. Ribot, J. H. 2013. Birds of Suriname. www.surinamebirds.nl/php/intro_e.php (accessed 1 November 2013). Sick, H. 1993. Birds in Brazil. Princeton Univ. Press. Skutch, A. F. 1966. Life history notes on three tropical American cuckoos. Wilson Bull. 78: 139-165. Tostain, O., Dujardin, J.-L., Érard, C. & Thiollay, J.-M. 1992. Oiseaux de Guyane. Société d’Études Ornithologiques, Brunoy. Addresses: Johan Ingels, Galgenberglaan 9, B-9070 Destelbergen, Belgium, e-mail: [email protected]. Mathias Fernandez, 2 rue Eugène Lony, F-97354 Rémire-Montjoly, France, e-mail: mathias.parare@ yahoo.fr